Anatomy of the coracoid and diversity of the Procellariiformes (Aves) in the Oligocene of Europe


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Abstract: Two European species of the Diomedeoididae, an extinct family of procellariiform (tube-nose) birds, have hitherto been distinguished primarily by size of their limb bones. Here, we describe an Early Oligocene (Rupelian) procellariifom coracoid that in all probability represents the larger species, Diomedeoides lipsiensis, and compare it to the coracoids of smaller diomedeoidids and extant procellariiforms. Using multivariate (Principal Component Analysis) and univariate analyses, we demonstrate that nearly all measurements are heavily size dependent, which makes the proportions and some other shape characters of little use as phylogenetic markers. Among eight measurements, the coracoid corpus width shows the highest correlation (higher than corpus depth) with body mass, permitting a precise calculation of over twofold difference in body mass between D. lipsiensis and smaller species. Among 16 qualitative characters analysed, the majority proved too variable to be used as markers of interfamily relationships and only 2–3, the ventral intermuscular line, sternocoracoid articulation (divided vs. undivided), and, with reservations, epimarginal crest vary consistently between the families. By far the most variable is the acrocoracoid process that tends to be deeper (more elongate dorsoventrally) in larger petrels but not in the albatrosses. However, the detailed shapes of the acrocoracoid heads are highly genus specific and suggest a genus-level diversity among the Diomedeoididae from the Oligocene of Europe. The common features of the diomedeoidid coracoids are best interpreted as plesiomorphies, which accounts for some similarities to the Oceanitinae (that are probably basal among the crown-group procellariiforms). The evidence from the coracoid is consistent with a stem-group position of the Diomedeoididae as previously proposed by others. We emphasize the need of a group-specific character analysis, primarily of allometries and levels of character variation, prior to a phylogenetic reconstruction.